Experimental data on photophysiology and growth rates of Pararotalia calcariformata

The eastern Mediterranean is a hotspot of biological invasions. Numerous species of Indo-pacific origin have colonized the Mediterranean in recent times, including tropical symbiont-bearing foraminifera. Among these is the species Pararotalia calcariformata. Unlike other invasive foraminifera, this species has been discovered only two decades ago and is restricted to the eastern Mediterranean coast. Combining ecological, genetic and physiological observations, we attempt to explain the recent invasion of this species in the Mediterranean Sea. Using morphological and genetic data, we confirm the species attribution to P. calcariformata McCulloch 1977 and identify its symbionts as a consortium of diatom species dominated by Minutocellus polymorphus. We document photosynthetic activity of its endosymbionts using Pulse Amplitude Modulated Fluorometry and test the effects of elevated temperatures on growth rates of asexual offspring. The culturing of asexual offspring for 120 days shows a 30-day period of rapid growth followed by a period of slower growth. A subsequent 48-day temperature sensitivity experiment indicates a similar developmental pathway and high growth rate at 28°C, whereas an almost complete inhibition of growth was observed at 20°C and 35°C. This indicates that the offspring of this species may have lower tolerance to cold temperatures than what would be expected for species native to the Mediterranean. We expand this hypothesis by applying a Species Distribution Model (SDM) based on modern occurrences in the Mediterranean using three environmental variables: irradiance, turbidity and yearly minimum temperature. The model reproduces the observed restricted distribution and indicates that the range of the species will drastically expand westwards under future global change scenarios. We conclude that P. calcariformata established a population in the Levant because of the recent warming in the region. In line with observations from other groups of organisms, our results indicate that continued warming of the eastern Mediterranean will facilitate the invasion of more tropical marine taxa into the Mediterranean, disturbing local biodiversity and ecosystem structure.

Benthic megafaunal composition at the Arctic deep-sea observatory HAUSGARTEN

The Long-Term Ecological Research (LTER) observatory HAUSGARTEN, in the eastern Fram Strait, provides us the valuable ability to study the composition of benthic megafaunal communities through the analysis of seafloor photographs. This, in combination with extensive sampling campaigns, which have yielded a unique data set on faunal, bacterial, biogeochemical and geological properties, as well as on hydrography and sedimentation patterns, allows us to address the question of why variations in megafaunal community structure and species distribution exist within regional (60-110 km) and local (<4 km) scales. Here, we present first results from the latitudinal HAUSGARTEN gradient, consisting of three different stations (N3, HG-IV, S3) between 78°30'N and 79°45'N (2351 - 2788 m depth), obtained via the analysis of images acquired by a towed camera (OFOS - Ocean Floor Observation System) in 2011. We assess variability in megafaunal densities, species composition and diversity as well as biotic and biogenic habitat features, which may cause the patterns observed. While there were significant regional-scale differences in megafaunal composition and densities between the stations (N3 = 26.74 ± 0.63; HG-IV = 11.21 ± 0.25; S3 = 18.34 ± 0.39 individuals/m**2), significant local differences were only found at HG-IV. Regional-scale variations may be due to the significant differences in ice coverage at each station as well as the different quantities of protein available, whereas local-scale differences at HG-IV may be a result of variation in bottom topography or factors not yet identified.

Table 1: Soil chemical properties of the analysed depth profiles in the Venice lagoon

Visual traces of iron reduction and oxidation are linked to the redox status of soils and have been used to characterise the quality of agricultural soils.We tested whether this feature could also be used to explain the spatial pattern of the natural vegetation of tidal habitats. If so, an easy assessment of the effect of rising sea level on tidal ecosystems would be possible. Our study was conducted at the salt marshes of the northern lagoon of Venice, which are strongly threatened by erosion and rising sea level and are part of the world heritage 'Venice and its lagoon'. We analysed the abundance of plant species at 255 sampling points along a land-sea gradient. In addition, we surveyed the redox morphology (presence/absence of red iron oxide mottles in the greyish topsoil horizons) of the soils and the presence of disturbances. We used indicator species analysis, correlation trees and multivariate regression trees to analyse relations between soil properties and plant species distribution. Plant species with known sensitivity to anaerobic conditions (e.g. Halimione portulacoides) were identified as indicators for oxic soils (showing iron oxide mottles within a greyish soil matrix). Plant species that tolerate a low redox potential (e.g. Spartina maritima) were identified as indicators for anoxic soils (greyish matrix without oxide mottles). Correlation trees and multivariate regression trees indicate the dominant role of the redox morphology of the soils in plant species distribution. In addition, the distance from the mainland and the presence of disturbances were identified as tree-splitting variables. The small-scale variation of oxygen availability plays a key role for the biodiversity of salt marsh ecosystems. Our results suggest that the redox morphology of salt marsh soils indicates the plant availability of oxygen. Thus, the consideration of this indicator may enable an understanding of the heterogeneity of biological processes in oxygen-limited systems and may be a sensitive and easy-to-use tool to assess human impacts on salt marsh ecosystems.

(Table 2) Planktic foraminifera and their diversity indices of sediment surface samples from the Atlantic Ocean

Species distribution patterns in planktonic foraminiferal assemblages are fundamental to the understanding of the determinants of their ecology. Until now, data used to identify such distribution patterns was mainly acquired using the standard >150 µm sieve size. However, given that assemblage shell size-range in planktonic foraminifera is not constant, this data acquisition practice could introduce artefacts in the distributional data. Here, we investigated the link between assemblage shell size-range and diversity in Recent planktonic foraminifera by analysing multiple sieve-size fractions in 12 samples spanning all bioprovinces of the Atlantic Ocean. Using five diversity indices covering various aspects of community structure, we found that counts from the >63 µm fraction in polar oceans and the >125 µm elsewhere sufficiently approximate maximum diversity in all Recent assemblages. Diversity values based on counts from the >150 µm fraction significantly underestimate maximum diversity in the polar and surprisingly also in the tropical provinces. Although the new methodology changes the shape of the diversity/sea-surface temperature (SST) relationship, its strength appears unaffected. Our analysis reveals that increasing diversity in planktonic foraminiferal assemblages is coupled with a progressive addition of larger species that have distinct, offset shell-size distributions. Thus, the previously documented increase in overall assemblage shell size-range towards lower latitudes is linked to an expanding shell-size disparity between species from the same locality. This observation supports the idea that diversity and shell size-range disparity in foraminiferal assemblages are the result of niche separation. Increasing SST leads to enhanced surface water stratification and results in vertical niche separation, which permits ecological specialisation. Specific deviations from the overall diversity and shell-size disparity latitudinal pattern are seen in regions of surface-water instability, indicating that coupled shell-size and diversity measurements could be used to reconstruct water column structures of past oceans.

Abundance and characteristics of Polychaeta taxa and associated sediment during POLARSTERN campaign ANT-XXII/3 (ANDEEP-IIII)

During the austral summer of 2005, the Weddell deep sea and adjacent basins were sampled in the course of the ANDEEP III project. In this study, 19 epibenthic-sledge stations are analyzed, with a focus on species diversity and distribution patterns of polychaetes. The polychaete fauna of the deep Southern Ocean has been found to be similarly speciose and diverse compared with deep-sea basins worldwide. Also, in depths below 2,000 m many polychaete species do not seem to be endemic for certain areas but are rather far spread within the Southern Ocean and beyond. Therefore, ongoing faunal exchanges between adjacent basins, even beyond the Antarctic convergence, are strongly suggested, ruling out a general isolation of the Southern Ocean deep-sea benthos. Driving forces behind species distribution patterns were investigated. The findings indicate that polychaete species' distribution in the Southern Ocean deep sea is rather dependent on local environment than depths.

Dinoflagellates of multicorer surface sediments

Only very few studies focus on recent calcareous dinoflagellate cyst diversity, geographic distribution and ecology, so that information on the distribution patterns and environmental affinities of individual cyst species is extremely limited. This information is, however, essential if we want to use calcareous dinoflagellate cysts for palaeoenvironmental reconstruction. Surface sediment samples from the generally oligotrophic western equatorial Atlantic Ocean, offshore northeast Brazil, were therefore quantitatively analysed for their calcareous dinoflagellate cyst content, including the calcareous vegetative coccoid Thoracosphaera heimii. Seven calcareous dinoflagellate cyst species/morphotypes and T. heimii were encountered in high concentrations throughout the area. Substantial differences in the distribution patterns were observed. The highest concentrations of cysts are found in sediments of the more oligotrophic, oceanic regions, beyond the influence of Amazon River discharge waters. Dinoflagellates producing calcareous cysts thus appear to be capable of surviving low nutrient concentrations and produce large numbers of cysts in relatively stable and predictable environments affected by minimal seasonality. To test for the environmental affinities of individual species, distribution patterns in surface sediments were compared with temperature, salinity, density and stratification gradients within the upper water column (0-100 m) over different times of the year, using principal components analysis and redundancy analysis. T. heimii and four of the seven encountered cyst species (Sphaerodinella? albatrosiana, two morphotypes of Sphaerodinella? tuberosa and Scrippsiella regalis) relate to these parameters significantly and the variations in the cyst associations appear to be associated with the different surface water currents characterising the area. The results imply that calcareous dinoflagellate cyst distributions can potentially be used to distinguish between different open oceanic environments and they could, therefore, be useful in tracing water mass movements throughout the late Quaternary.

Ostracode relative abundances and geochemistry in surface sediments from Lake Ohrid

Understanding the ecology of bioindicators such as ostracods is essential in order to reconstruct past environmental and climate change from analysis of fossil assemblages preserved in lake sediment cores. Knowledge of the ecology of ancient Lake Ohrid's ostracod fauna is very limited and open to debate. In advance of the Ohrid ICDP-Drilling project, which has potential to generate high-resolution long-term paleoenvironmental data of global importance in paleoclimate research, we sampled Lake Ohrid and a wide range of habitat types in its surroundings to assess 1) the composition of ostracod assemblages in lakes, springs, streams, and short-lived seasonal water bodies, 2) the geographical distribution of ostracods, and 3) the ecological characteristics of individual ostracod species. In total, 40 species were collected alive, and seven species were preserved as valves and empty carapaces. Of the 40 ostracod species, twelve were endemic to Lake Ohrid. The most common genus in the lake was Candona, represented by 13 living species, followed by Paralimnocythere, represented by five living species. The most frequent species was Cypria obliqua. Species with distinct distributions included Heterocypris incongruens, Candonopsis kingsleii, and Cypria lacustris. The most common species in shallow, flooded areas was H. incongruens, and the most prominent species in ditches was C. kingsleii. C. lacustris was widely distributed in channels, springs, lakes, and rivers. Statistical analyses were performed on a "Lake Ohrid" dataset, comprising the subset of samples from Lake Ohrid alone, and an "entire" dataset comprising all samples collected. The unweighted pair group mean average (UPGMA) clustering was mainly controlled by species-specific depth preferences. Canonical Correspondence Analysis (CCA) with forward selection identified water depth, water temperature, and pH as variables that best explained the ostracod distribution in Lake Ohrid. The lack of significance of conductivity and dissolved oxygen in CCA of Ohrid data highlight the uniformity across the lake of the well-mixed waters. In the entire area, CCA revealed that ostracod distribution was best explained by water depth, salinity, conductivity, pH, and dissolved oxygen. Salinity was probably selected by CCA due to the presence of Eucypris virens and Bradleystrandesia reticulata in short-lived seasonal water bodies. Water depth is an important, although indirect, influence on ostracod species distribution which is probably associated with other factors such as sediment texture and food supply. Some species appeared to be indicators for multiple environmental variables, such as lake level and water temperature.

Diatom abundance in surface sediments of the Southern Ocean

Based on the quantitative analysis of diatom assemblages preserved in 274 surface sediment samples recovered in the Pacific, Atlantic and western Indian sectors of the Southern Ocean we have defined a new reference database for quantitative estimation of late-middle Pleistocene Antarctic sea ice fields using the transfer function technique. The Detrended Canonical Analysis (DCA) of the diatom data set points to a unimodal distribution of the diatom assemblages. Canonical Correspondence Analysis (CCA) indicates that winter sea ice (WSI) but also summer sea surface temperature (SSST) represent the most prominent environmental variables that control the spatial species distribution. To test the applicability of transfer functions for sea ice reconstruction in terms of concentration and occurrence probability we applied four different methods, the Imbrie and Kipp Method (IKM), the Modern Analog Technique (MAT), Weighted Averaging (WA), and Weighted Averaging Partial Least Squares (WAPLS), using logarithm-transformed diatom data and satellite-derived (1981-2010) sea ice data as a reference. The best performance for IKM results was obtained using a subset of 172 samples with 28 diatom taxa/taxa groups, quadratic regression and a three-factor model (IKM-D172/28/3q) resulting in root mean square errors of prediction (RMSEP) of 7.27% and 11.4% for WSI and summer sea ice (SSI) concentration, respectively. MAT estimates were calculated with different numbers of analogs (4, 6) using a 274-sample/28-taxa reference data set (MAT-D274/28/4an, -6an) resulting in RMSEP's ranging from 5.52% (4an) to 5.91% (6an) for WSI as well as 8.93% (4an) to 9.05% (6an) for SSI. WA and WAPLS performed less well with the D274 data set, compared to MAT, achieving WSI concentration RMSEP's of 9.91% with WA and 11.29% with WAPLS, recommending the use of IKM and MAT. The application of IKM and MAT to surface sediment data revealed strong relations to the satellite-derived winter and summer sea ice field. Sea ice reconstructions performed on an Atlantic- and a Pacific Southern Ocean sediment core, both documenting sea ice variability over the past 150,000 years (MIS 1 - MIS 6), resulted in similar glacial/interglacial trends of IKM and MAT-based sea-ice estimates. On the average, however, IKM estimates display smaller WSI and slightly higher SSI concentration and probability at lower variability in comparison with MAT. This pattern is a result of different estimation techniques with integration of WSI and SSI signals in one single factor assemblage by applying IKM and selecting specific single samples, thus keeping close to the original diatom database and included variability, by MAT. In contrast to the estimation of WSI, reconstructions of past SSI variability remains weaker. Combined with diatom-based estimates, the abundance and flux pattern of biogenic opal represents an additional indication for the WSI and SSI extent.

Planktic foraminifera analyses of surface sediments and satellite data in the Canary Islands region

The Canary Islands region occupies a key position with respect to biogeochemical cycles, with the zonal transition from oligotrophic to nutrient-rich waters and the contribution of Saharan dust to the particle flux. We present the distribution of geochemical proxies (TOC, carbonate, d15N, d13Corg, C/N-ratio) and micropaleontological parameters (diatoms, dinoflagellates, foraminifera, pteropods), in 80 surface-sediment samples in order to characterise the influence of coastally upwelled water on the domain of the subtropical gyre. Results of the surface-sediment analyses confirmed the high biomass gradient from the coast to the open ocean inferred from satellite data of surface chlorophyll or SST. The distribution of total dinoflagellate cysts, the planktic foraminifera species Globigerina bulloides, the diatom resting spore Chaetoceros spp., and TOC concentration coincided well with the areas of strong filament production off Cape Ghir and Cape Yubi. The warm-water planktic foraminifera Globigerinoides ruber (white), the diatom Nitzschia spp., and the d15N-values showed the opposite trend with high values in the open ocean. Factor analyses on the planktic foraminifera species distribution indicated three major assemblages in the Canary Islands region that represent the present surface-water conditions from the upwelling influenced region via a mixing area towards the subtropical gyre.